Axolotl
Axolotl | |
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In the National Aquarium in Washington, D.C. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Amphibia |
Order: | Urodela |
Family: | Ambystomatidae |
Genus: | Ambystoma |
Species: | A. mexicanum
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Binomial name | |
Ambystoma mexicanum | |
IUCN range of the Axolotl.
Axolotl (Ambystoma mexicanum)
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Synonyms[3] | |
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The axolotl (
As of 2020[update], the axolotl was near
Axolotls should not be confused with the larval stage of the closely related tiger salamander (A. tigrinum), which are widespread in much of North America and occasionally become paedomorphic. Neither should they be confused with
Description
A sexually mature adult axolotl, at age 18–27 months, ranges in length from 15 to 45 cm (6 to 18 in), although a size close to 23 cm (9 in) is most common and greater than 30 cm (12 in) is rare. Axolotls possess features typical of salamander larvae, including external gills and a caudal fin extending from behind the head to the vent.[13][14] External gills are usually lost when salamander species mature into adulthood, although the axolotl maintains this feature.[15] This is due to their neoteny evolution, where axolotls are much more aquatic than other salamander species.[16]
Their heads are wide, and their eyes are lidless. Their limbs are underdeveloped and possess long, thin digits. Males are identified by their swollen
Axolotls have barely visible
Axolotls have four pigmentation genes; when mutated, they create different color variants. The normal wild-type animal is brown or tan with gold speckles and an olive undertone. The five most common mutant colors are listed below.
- Leucistic: pale pink with black eyes.
- Xanthic: grey, with black eyes.
- Albino: pale pink or white, with red eyes, which is more common in axolotls than other species.
- Melanoid: all black or dark blue with no gold speckling or olive tone.
In addition, there is wide individual variability in the size, frequency, and intensity of the gold speckling, and at least one variant develops a black and white piebald appearance upon reaching maturity. [17] Because pet breeders frequently cross the variant colors, double homozygous mutants are common in the pet trade, especially white/pink animals with pink eyes that are double homozygous mutants for both the albino and leucistic trait.[18] Axolotls also have some limited ability to alter their color to provide better camouflage by changing the relative size and thickness of their melanophores.[19]
Habitat and ecology
The axolotl is native only to the freshwater of Lake Xochimilco and Lake Chalco in the Valley of Mexico. Lake Chalco no longer exists, having been drained as a flood control measure, and Lake Xochimilco remains a remnant of its former self, existing mainly as canals. The water temperature in Xochimilco rarely rises above 20 °C (68 °F), although it may fall to 6–7 °C (43–45 °F) in the winter, and perhaps lower.[20]
Surveys in 1998, 2003, and 2008 found 6,000, 1,000, and 100 axolotls per square kilometer in its Lake Xochimilco habitat, respectively.[21] A four-month-long search in 2013, however, turned up no surviving individuals in the wild. Just a month later, two wild ones were spotted in a network of canals leading from Xochimilco.[22]
The wild population has been put under heavy pressure by the growth of Mexico City. The axolotl is currently on the International Union for Conservation of Nature's annual Red List of threatened species. Non-native fish, such as African tilapia and Asian carp, have also recently been introduced to the waters. These new fish have been eating the axolotls' young, as well as their primary source of food.[23]
Axolotls are members of the tiger salamander, or
Diet
The axolotl is carnivorous, consuming small prey such as mollusks,[24] worms, insects, other arthropods,[24] and small fish in the wild. Axolotls locate food by smell, and will "snap" at any potential meal, sucking the food into their stomachs with vacuum force.[25]
Use as a model organism
Today, the axolotl is still used in research as a
Regeneration
The feature of the axolotl that attracts most attention is its healing ability: the axolotl does not heal by scarring and is capable of the regeneration of entire lost appendages in a period of months, and, in certain cases, more vital structures, such as tail, limb, central nervous system, and tissues of the eye and heart.[28] They can even restore less vital parts of their brains. They can also readily accept transplants from other individuals, including eyes and parts of the brain—restoring these alien organs to full functionality. In some cases, axolotls have been known to repair a damaged limb, as well as regenerating an additional one, ending up with an extra appendage that makes them attractive to pet owners as a novelty. In metamorphosed individuals, however, the ability to regenerate is greatly diminished. The axolotl is therefore used as a model for the development of limbs in vertebrates.[29] There are three basic requirements for regeneration of the limb: the wound epithelium, nerve signaling, and the presence of cells from the different limb axes.[30] A wound epidermis is quickly formed by the cells to cover up the site of the wound. In the following days, the cells of the wound epidermis divide and grow quickly forming a blastema, which means the wound is ready to heal and undergo patterning to form the new limb.
It is believed that during limb generation, axolotls have a different system to regulate their internal macrophage level and suppress inflammation, as scarring prevents proper healing and regeneration.[31] However, this belief has been questioned by other studies.[32] The axolotl's regenerative properties leave the species as the perfect model to study the process of stem cells and its own neoteny feature. Current research can record specific examples of these regenerative properties through tracking cell fates and behaviors, lineage tracing skin triploid cell grafts, pigmentation imaging, electroporation, tissue clearing and lineage tracing from dye labeling. The newer technologies of germline modification and transgenesis are better suited for live imaging the regenerative processes that occur for axolotls.[33]
Genome
The 32 billion
Neoteny
Most amphibians begin their lives as
The genes responsible for neoteny in laboratory animals may have been identified; however, they are not linked in wild populations, suggesting
Metamorphosis
The axolotl's body has the capacity to go through metamorphosis if given the necessary hormone, but axolotls do not produce it, and must be exposed to it from an external source, after which an axolotl undergoes an artificially-induced metamorphosis and begins living on land.
Role of iodine
In animals with functioning thyroid glands, iodine in the form of iodide is selectively gathered into the colloid of the thyroud. Inside the colloid, iodide is reduced to elemental iodine (I2), which reacts with the
In the absence of induced metamorphosis, larval axolotls start absorbing iodide into their thyroid glands at 30 days postfertilization. Larval axolotls do produce thyroid hormone from iodide, but the amount appears highly variable. Adult axolotls do not produce thyroid hormone unless metamorphism is triggered.[40]
An axolotl undergoing metamorphosis experiences a number of physiological changes that help them adapt to life on land. These include increased muscle tone in limbs, the absorption of gills and fins into the body, the development of eyelids, and a reduction in the skin's permeability to water, allowing the axolotl to stay more easily hydrated when on land. The lungs of an axolotl, though present alongside gills after reaching non-metamorphosed adulthood, develop further during metamorphosis.[45]
An axolotl that has gone through metamorphosis resembles an adult plateau tiger salamander, though the axolotl differs in its longer toes.[citation needed] Among hobbyists, the process of artificially inducing metamorphosis can often result in death during or even following a successful attempt, and so casual hobbyists are generally discouraged from attempting to induce metamorphosis in pet axolotls.[45] Morphed pet axolotls should be given solid footholds in their enclosure to satisfy their need for land. They should not be given live animals as food.[46]
History
Six adult axolotls (including a leucistic specimen) were shipped from
In other salamanders
Many other species within the axolotl's genus are also either entirely neotenic or have neotenic populations.
Neoteny has been observed in all
Threats
Axolotls are only native to the Mexican Central Valley. Although the native axolotl population once extended through most of the lakes and wetlands that make up this region, the native habitat is now limited to Lake Xochimilco as a result of the expansion of Mexico City. Lake Xochimilco is not a large body of water, but rather a small series of artificial channels, small lakes, and temporary wetlands.
Lake Xochimilco has poor water quality, caused by the region's aquaculture and agriculture demands. It is also maintained by inputs of only partially treated wastewater. Water quality tests reveal a low nitrogen-phosphorus ratio and a high concentration of chlorophyll a, which are indicative of an oxygen-poor environment that is not well-suited for axolotls.[49] In addition, the intensive use of pesticides from agriculture around Lake Xochimilco causes run off into the lake and a reduction of habitat quality for axolotls. The pesticides used contain chemical compounds that studies show to sharply increase mortality in axolotl embryos and larvae. Of the surviving embryo and larvae, there is also an increase of morphological, behavior, and activity abnormalities.[50]
Another factor that threatens the native axolotl population is the introduction of invasive species such as the Nile tilapia and common carp. These invasive fish species threaten axolotl populations by eating their eggs or young and by out-competing them for natural resources. The presence of these species has also been shown to change the behavior of axolotls, causing them to be less active to avoid predation. This reduction in activity greatly impacts the axolotls foraging and mating opportunities.[51]
With such a small native population, there is a large loss of genetic diversity. This lack of genetic diversity can be dangerous for the remaining population, causing an increase in inbreeding and a decrease in general fitness and adaptive potential. It ultimately raises the axolotl's risk for extinction, something that they are already in danger of. Studies have found indicators of a low interpopulation gene flow and higher rates of genetic drift. These are likely the result of multiple “bottleneck” incidents in which events that kill off several individuals of a population occur and sharply reduce the genetic diversity of the remaining population. The offspring produced after bottleneck events have a greater risk of showing decreased fitness and are often less capable of adaptation down the line. Multiple bottleneck events can have disastrous effects on a population. Studies have also found high rates of relatedness that are indicative of inbreeding. Inbreeding can be especially harmful as it can cause an increase in the presence of deleterious, or harmful, genes within a population.[52] The detection of introgressed tiger salamander (A. tigrinum) DNA in the laboratory axolotl population raises further concerns about the suitability of the captive population as an ark for potential reintroduction purposes.[53]
There has been little improvement in the conditions of the lake or the population of native axolotls. Many scientists are focusing their conservation efforts on translocation of captive-bred individuals into new habitats or reintroduction into Lake Xochimilco. The Laboratorio de Restauracion Ecologica (LRE) in the Universidad Nacional Autonoma de Mexico (UNAM) has built up a population of more than 100 captive-bred individuals. These axolotls are mostly used for research by the lab but plans of a semi-artificial wetland inside the university have been established and the goal is to establish a viable population of axolotls within it. Studies have shown that captive-bred axolotls that are raised in a semi-natural environment can catch prey, survive in the wild, and have moderate success in escaping predators. These captive-bred individuals can be introduced into unpolluted bodies of water or back into Lake Xochimilco to establish or re-establish a wild population.[54][55]
Captive care
Salts, such as Holtfreter's solution, are often added to the water to prevent infection.[60]
In captivity, axolotls eat a variety of readily available foods, including trout and salmon pellets, frozen or live
Substrates are another important consideration for captive axolotls, as axolotls (like other amphibians and reptiles) tend to ingest bedding material together with food[62] and are commonly prone to gastrointestinal obstruction and foreign body ingestion.[63] Some common substrates used for animal enclosures can be harmful for amphibians and reptiles. Gravel (common in aquarium use) should not be used, and is recommended that any sand consists of smooth particles with a grain size of under 1mm.[62] One guide to axolotl care for laboratories notes that bowel obstructions are a common cause of death, and recommends that no items with a diameter below 3 cm (or approximately the size of the animal's head) should be available to the animal.[64]
There is some evidence that axolotls might seek out appropriately-sized gravel for use as
Cultural significance
The species is named after the
They appear in the works of Mexican muralist
The
See also
- Mudpuppies
- Olm
- Texas salamander
- Texas blind salamander
- Lake Patzcuaro salamander
- Barred tiger salamander
- Amphibious fish
- Handfish
- Regenerative biomedicine
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External links
- Ambystomatidae at Curlie
- Follow the Eggs, Hatchlings and Juveniles
- Mating Dance and Laying Eggs
- Follow the Eggs and Hatchlings (2nd Batch)
- Indiana U Axolotl Colony
- University of KY Axolotl Colony
- Mystical amphibian venerated by Aztecs nears extinction
- The animal that’s everywhere and nowhere
- Encyclopædia Britannica. Vol. 20 (11th ed.). 1911. p. 63. .